Genetic Divergence of Native Palms of Oenocarpus Distichus Considering
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www.nature.com/scientificreports OPEN Genetic divergence of native palms of Oenocarpus distichus considering biometric fruit variables Received: 9 August 2018 Gleidson Guilherme Caldas Mendes1, Mônica Trindade Abreu de Gusmão2, Accepted: 1 March 2019 Thales Guilherme Vaz Martins1, Renato Domiciano Silva Rosado 3, Published: xx xx xxxx Rayanne Savina Alencar Sobrinho2, Andrei Caíque Pires Nunes4, Wellington Souto Ribeiro5 & José Cola Zanuncio6 Oenocarpus distichus presents economic, ecological, and dietary potential for pulp market processed in natura. Germplasm conservation and genetic improvement depend on genetic divergence studies. The objective was to quantify genetic divergence in a native population of O. distichus genotypes based on fruit biometrics. The fruit length and width, fruit mass, pulp and seed, pulp and almond thickness, and pulp yield per fruit were evaluated. All fruit biometric characteristics of O. distichus palms show genetic variability. Genetic variations among genotypes are essential for predicting heredity and heterosis, which are essential for improving O. distichus production. Pulp yield and seed mass were negatively correlated. Almond thickness and pulp, seed mass and transverse diameter were positively correlated. Genetic distances between pair of genotypes ranged from 0.07 to 48.10 with three genetically distinct groups. The seed mass, almond thickness and transverse diameter contributed to genetic divergence. Heritability estimates the genetic control that can be obtained from O. distichus germplasm. Correlations between the variable pair reduce the evaluation efort and the resources to measure the genotype allocations in heterogeneous groups presenting high genetic variability. This makes it possible to select individuals for hybridization programs with F1 generation gains. Correlation and relative contribution networks, based on relationships graphical between fruit biometric characteristics, allow the variables selection with less efort and fewer measurements. O. distichus fruit biometric characters are efcient to quantify genetic divergence between genotypes. Te genetic diversity of Brazilian Amazonian plants is signifcant but remains poorly understood. Te sites iden- tifcation for conservation and germplasm collection in this region is a great challenge, especially for some native palm trees1. Palm fruits and palm hearts native to the Amazon are consumed by local communities but are poorly known at the national and international levels2. Oenocarpus spp. plants have economic, ecological and dietary potential and are exploited through extraction by Amazonian communities1,3. Six species of the genus Oenocarpus spp. are native to Brazil, but not endemic, including O. distichus Mart., O. bacaba Mart., O. minor Mart. and O. mapora H. Karten and O. pataua Mart.4. Oenocarpus distichus (also known as bacaba-de-leque) has high potential in the palm and processed pulp markets, including its use in the produc- tion of “bacaba wine”, a nutritious energy drink with commercial potential similar to that of Euterpe oleracea Mart.5. Oenocarpus distichus phytochemicals have phenolic compounds correlated with antioxidant capacity2,6,7 and vitamin E, ribofavin, insoluble fber, energy, and minerals, especially potassium and calcium8. Tis plant has oil 1Departamento de Engenharia Florestal, Universidade Federal de Viçosa, 36570-900, Viçosa, Minas Gerais, Brazil. 2Instituto de Ciências Agrárias, Universidade Federal Rural da Amazônia, 66077-030, Belém, Pará, Brazil. 3Departamento de Estatística Aplicada e Biometria, Universidade Federal de Viçosa, 36570-900, Viçosa, Minas Gerais, Brazil. 4Centro de Formação em Ciências Agroforestais, Universidade Federal do Sul da Bahia, 45613-204, Itabuna, Bahia, Brazil. 5Programa de Pós-graduação em Horticultura Tropical, Universidade Federal de Campina Grande, 8, Rua Jairo Vieira Feitosa, 58840-000, Pombal, Paraíba, Brazil. 6Departamento de Entomologia/BIOAGRO, Universidade Federal de Viçosa, 36570-900, Viçosa, Minas Gerais, Brazil. Correspondence and requests for materials should be addressed to G.G.C.M. (email: [email protected]) SCIENTIFIC REPORTS | (2019) 9:4943 | https://doi.org/10.1038/s41598-019-41507-4 1 www.nature.com/scientificreports/ www.nature.com/scientificreports Characters GMS RMS Mean CVe (%) CVg% CVg/CVe h²mp FL (mm) 13.45** 1.20 16.40 6.67 5.70 0.85 91.09 WF (mm) 15.16** 0.99 14.76 6.76 6.82 1.01 93.45 FM (g) 2.60** 0.08 2.54 11.16 16.72 1.50 96.82 PM (g) 0.40** 0.02 0.86 15.09 19.27 1.28 95.81 SM (g) 1.30** 0.06 1.68 14.23 17.75 1.25 95.61 PT (mm) 0.25** 0.04 1.65 11.56 7.47 0.65 85.38 AT (mm) 3.06** 0.19 5.53 7.86 8.19 0.99 93.82 PY (%) 164.91** 18.99 34.06 12.80 9.48 0.74 88.48 Table 1. Variance analysis and genetic parameters estimates for morphological characters of Oenocarpus distichus fruits. FL- fruit length; WF- fruit width, FM- fruit mass; PM- pulp mass; SM- seed mass; PT- pulp thickness; AT- almond thickness; PY- pulp yield per fruit; GMS- genotype mean square; RMS- residue mean square; CVe- experimental variation coefcient; CVg- genetic variation coefcient; h²mp- mean progeny heritability level; **Signifcant at 1% by F test. with organoleptic properties similar to those of olive oil and raw material for the food industry with proteins of high biological value (40% more than soy)5 and adequate to manufacture ice cream and soap9. Te genetic diversity in O. distichus native populations, from the morphological characteristics of their fruits, needs to be better understood for germplasm inclusion. Tis is necessary for breeding programs to conserve native populations in extractive areas. Oenocarpus distichus genetic improvement and conservation programs depend on fruits biometric characterization and genetic parameter studies of genotypes showing promise for germplasm banks and conservation programs of this plant4,10. Multivariate analyses allow selecting individuals for breeding programs and morphological or molecular char- acteristic identifcation to evaluate the divergence of native and exotic plants such as E. oleracea11, Teobroma grandiforum Willd. Ex Spreng.12, Elaeis guineenses Jacq.13 and Phoenix dactylifera L.14 in forest fragments15. Grouping methods, based on prefxed similarity/dissimilarity measures, can identify divergent groups. Group determination, based on a standard, universally, accepted method does not exist, but individuals of the same group should be as homogeneous as possible and diferent from the others16. Fruit characteristics are important for characterization, individual selection and genotypes choice during recombination cycles15,17. Genetic divergence, important for prospecting in plant breeding programs, allows us to know the available germplasm to evaluate similar or divergent groups to identify hybrid combinations with greater heterozygosity and heterotrophic efect18. Genetic divergence will allow us to know the available ger- mplasm to predict combinations with higher heterozygosity so that there will be more possibilities of recovering fruitful Oenocarpus distichus genotypes in the segregant generations. Distribution among groups will allows the early determination of the best combinations and reduces the unneeded crosses number, prioritizing more con- trasting groups for obtention of combination with higher heterosis. Biometric fruit variables diferences such as morphological and production are regarded in the dissimilarity quantifcation. Tose genetic variations among Oenocarpus distichus genotypes are essential for predicting heredity and level of heterosis, which are essential for improving Oenocarpus distichus production. Te objective of the present study was to quantify the genetic divergence between Oenocarpus distichus gen- otypes in a native population in the northeastern region of Pará, Amazon rainforest, Brazil, based on fruit biom- etric characteristics. Results All O. distichus fruit biometric characteristics (F-test < 0.01) show genetic variability (Table 1) with residual var- iation coefcients (CVe) of 6.67% for fruit length (FL) at 15.09% for pulp mass (PM), indicating high experimen- tal precision. Te genetic variation coefcient (CVg) values, mainly of the PM, seed mass (SM) and fruit mass (FM) characters, confrmed the genetic variation between the O. distichus genotypes. Te relationship between CVg/CVe was greater than 1 for the characteristics fruit width (WF), FM, PM and SM. Matrix inheritance esti- 2 mates for average ones (h mp) ranged from 85.38 for pulp thickness (PT) to 96.81 for FM. Te mean values of biometric characteristics of O. distichus fruits and Scott-Knott test at 5% probability are given in supplementary information. Te biometry correlation matrix of the O. distichus fruits indicate estimates for the last two eigenvalues near to zero, resulting in two linear relationships determining the harmful efects of multicollinearity (Table 2). Te condition number for this matrix indicated NC > 1000, that is, severe colinearity. Te eigenvectors, associated with the last two eigenvalues destacados, indicated that the FM and PM characteristics (highest absolute values highlighted in bold) are responsible for multicollinearity. Te new NC matrix lower than 100, afer the exclusion of the FM and PM variables, indicates weak collinearity (Table 3). In addition to the CN, small single values were observed for FM (0.0503) and PM (0.0043) and high condition index FM (46.60) and PM (537.90), indicating serious problems caused by multicollinearity. VIF values can also be used to detect the existence